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 Environ. Res. Lett. 13 (2018) 024017 https://doi.org/10.1088/1748-9326/aaa273

LETTER

Feed conversion efficiency in aquaculture: do we

OPEN ACCESS
measure it correctly?
RECEIVED
9 August 2017
Jillian P Fry1,2,3,6 , Nicholas A Mailloux1 , David C Love1,2 , Michael C Milli1 and Ling Cao4,5
REVISED 1 Johns Hopkins Center for a Livable Future, Johns Hopkins University, 615 N Wolfe Street, Baltimore, MD, United States of America
13 December 2017 2 Department of Environmental Health and Engineering, Bloomberg School of Public Health, Johns Hopkins University, 615 N Wolfe
ACCEPTED FOR PUBLICATION
Street, Baltimore, MD, United States of America
18 December 2017 3 Department of Health, Behavior and Society, Bloomberg School of Public Health, Johns Hopkins University, 624 N Broadway,
PUBLISHED Baltimore, MD, United States of America
6 February 2018 4 Center on Food Security and the Environment, Stanford University, 616 Serra St, Stanford, CA, United States of America
5 Institute of Oceanography, Shanghai Jiao Tong University, Shanghai, People’s Republic of China
6 Author to whom any correspondence should be addressed.
Original content from
this work may be used
under the terms of the E-mail: jfry3@jhu.edu
Creative Commons
Attribution 3.0 licence. Keywords: aquaculture, agriculture, food security, animal production efficiency
Any further distribution
of this work must
Supplementary material for this article is available online
maintain attribution to
the author(s) and the
title of the work, journal Abstract
citation and DOI.
Globally, demand for food animal products is rising. At the same time, we face mounting, related
pressures including limited natural resources, negative environmental externalities, climate
disruption, and population growth. Governments and other stakeholders are seeking strategies to
boost food production efficiency and food system resiliency, and aquaculture (farmed seafood) is
commonly viewed as having a major role in improving global food security based on longstanding
measures of animal production efficiency. The most widely used measurement is called the ‘feed
conversion ratio’ (FCR), which is the weight of feed administered over the lifetime of an animal
divided by weight gained. By this measure, fed aquaculture and chickens are similarly efficient at
converting feed into animal biomass, and both are more efficient compared to pigs and cattle. FCR
does not account for differences in feed content, edible portion of an animal, or nutritional quality of
the final product. Given these limitations, we searched the literature for alternative efficiency
measures and identified ‘nutrient retention’, which can be used to compare protein and calories in
feed (inputs) and edible portions of animals (outputs). Protein and calorie retention have not been
calculated for most aquaculture species. Focusing on commercial production, we collected data on
feed composition, feed conversion ratios, edible portions (i.e. yield), and nutritional content of edible
flesh for nine aquatic and three terrestrial farmed animal species. We estimate that 19% of protein and
10% of calories in feed for aquatic species are ultimately made available in the human food supply,
with significant variation between species. Comparing all terrestrial and aquatic animals in the study,
chickens are most efficient using these measures, followed by Atlantic salmon. Despite lower FCRs in
aquaculture, protein and calorie retention for aquaculture production is comparable to livestock
production. This is, in part, due to farmed fish and shrimp requiring higher levels of protein and
calories in feed compared to chickens, pigs, and cattle. Strategies to address global food security
should consider these alternative efficiency measures.

1. Introduction [1, 2]. This is, in part, because an increasingly affluent

The global food system is a major force driving human-
ity towards bypassing multiple planetary boundaries,
including freshwater use, land use change, biodiver-
sity loss, climate change, and water quality degradation
and growing global human population is consuming
more meat and dairy products [3–6]. Food animal
products provide a concentrated source of calories,
protein, and some micronutrients. There are, however,
well-documented inefficiencies in terrestrial livestock

© 2018 The Author(s). Published by IOP Publishing Ltd

Environ. Res. Lett. 13 (2018) 024017
Giant tiger prawn
Common carp
Pangas catfish
Tilapia
Grass carp
Channel catfish
Whiteleg shrimp
Rainbow trout
Atlantic salmon
Aquaculture weighted avg.
Beef cattle
Pigs
Chicken
0 1
Figure 1. Feed conversion ratios for selected aquatic and terrestrial farmed animal species. Dots represent means and bars indicate
range. Lower values signify higher efficiency. Sources: Tacon and Metian (2008) [12], Smil (2013) [13], Shike (2013) [14], Zuidhof et
al (2014) [15], and Rabobank Research (2015) [16].
production. Approximately 36% of global crop-based
calories (3.41 × 1015 kcal) are fed to livestock, and of
those, just 12% enter the human food supply [7].
Aquaculture, or farmed seafood, is the fastest grow-
ing food animal sector and now contributes more
to the human food supply (by weight) than wild-
caught seafood (adjusting for wild-caught fish not
eaten by people) or beef [8, 9]. (We use the term,
seafood, to refer to aquatic animals caught or farmed
for human consumption in marine and freshwater set-
tings.) Seafood, from farmed and wild sources, provides
17% of global animal protein, and accounts for over
half of animal protein supplies in some developing
countries [8]. Aquaculture is heterogeneous in terms
of farmed species and production methods. Fed aqua-
culture, including both intensive and semi-intensive
systems, involves relatively high stocking densities and
either farm-made feeds or commercial compound feeds
formulated to meet nutritional requirements. Unfed
aquaculture includes filter-feeding molluscan shellfish
(e.g. oysters, clams, mussels) and aquatic plants (e.g.
microalgae, seaweed). Globally, aquaculture produc-
tion continues to expand and intensify. About 70% of
global aquaculture (excluding aquatic plants) relies on
commercial compound feed, and demand for commer-
cial feed is growing faster than the industry as a whole.
A significant proportion of aquaculture feed con-
tains ingredients made from wild-caught fish [10]. To
reduce pressure on depleted wild fisheries, the industry
is increasingly relying on alternative feed ingredients
including crop-based ingredients (e.g. soy, rapeseed,
wheat, groundnuts, and corn) and terrestrial animal
byproducts as substitutes for fishmeal and fish oil [11].
The efficiency with which animals turn feed into
meat and other food products, such as eggs or milk,
varies by species and production method. A common
2
2 3 4 5 6 7 8 9 10 11
Feed conversion ratio (kg of feed/kg weight gain)
measure of this efficiency is the feed conversion ratio
(FCR), calculated as the ratio of feed intake to weight
gain. Typical FCRs for animals raised using commer-
cial feeds and intensive production methods (i.e. not
extensive production like grazing) are as follows: beef
cattle: 6.0–10.0, pigs: 2.7–5.0, chickens: 1.7–2.0, and
farmed fish and shrimp: 1.0–2.4 (figure 1) [12–16].
Aquatic animals have lower (more efficient) FCRs than
large terrestrial animals in part because they expend
less energy to move, stay upright, and regulate their
body temperatures due to buoyancy and because most
are ectothermic [17, 18]. Expanding aquaculture is thus
widely viewed as an opportunity to meet rising demand
for animal products using less feed, especially compared
to pigs and cattle [19, 20]. FCR is a limited measure of
efficiency, however, because it only accounts for the
weight of feed inputs and not the nutritional content of
the feed, the portion of the animal that is inedible, or
the nutritional quality of the final product. Using FCRs
relies on an implicit assumption that various species
are similar across these areas, making FCR a potentially
flawed tool for cross-species comparisons.
We reviewed the literature and identified 13 differ-
ent approaches to measure aquatic animal production
efficiency beyond FCR (supplementary table S1 avail-
able at stacks.iop.org/ERL/13/024017/mmedia). Based
on our review, a more precise measure than FCR is
the efficiency with which an animal converts nutri-
ents in feed into nutrients for the human food supply,
specific examples of ‘nutrient retention’ measures are
sometimes called a ‘protein/calorie efficiency ratio’ or
‘protein/calorie retention’. These have been calculated
for major livestock products [7, 21] and for two aqua-
culture species (for example, see [22, 23]), but more
work is needed. Ytrestøyl et al calculated protein and
calorie efficiency for farmed salmon in Norway [22].
Environ. Res. Lett. 13 (2018) 024017

Table 1. Data used to calculate protein and calorie retention for selected aquatic and terrestrial farmed animal species.

Species FCRa Edible portion Feed contentc Human nutritiond

of animalb (g or kcal per 100 g of feed) (g or kcal per 100 g serving)
Protein Calories Protein Calories
Carps 1.5–2.0
Common carp – 0.36–0.54 17–45 175.8–554.2 18 109–127
Grass carp – 0.36–0.54 25 326.0–345.5 17–18 112–127
Catfishes 1.2–2.2
Channel catfish – 0.35–0.63 28–32 345–390 15–17 117–119
Pangas catfish – 0.35–0.63 26–32 339–388 15 97
Salmonids –
Atlantic salmon 1.2–1.5 0.58–0.88 35.5–44 372–554.5 20 208
Rainbow trout 1.0–2.0 0.40–0.82 40–47 383–454 20 141
Shrimps 1.2–2.4
Giant tiger prawn – 0.40 25–45 225–433 20 85
Whiteleg shrimp – 0.62–0.65 25–45 277–417 20 85
Tilapias 1.4–2.4 0.37–0.45 20–32 216–404.4 20 96
Cattle 6.0–10 0.52–0.64 7–15.4 188–339 15–20 214–276
Chicken 1.7–2.0 0.70–0.78 18–23 320 18.6 215
Pigs 2.7–5.0 0.68–0.76 13.2–20.9 326.5–335.1 15–18.2 211–304
a Data sources: Tacon and Metian (2008) (aquatic species) [12]; Smil (2013) (livestock species) [13]; Shike (2013) (cattle) [14]; Zuidhof et al

(2014) (chicken) [15]; Rabobank Research (2015) (pigs) [16].

b Data sources: see table S4.
c Data sources: see table S5.
d Data sources: USDA National Nutrient Database for Standard Reference [27]; Shauhua Zahn, Nanyang Technical University (personal

communication); Seafood Health Facts [28]; USDA National Nutrient Database terms used for beef: ‘composite of trimmed retail cuts,
separable lean and fat, trimmed to 1/8′′ fat, all grades, raw’ and ‘variety meats and by-products, mechanically separated beef, raw’; USDA
National Nutrient Database term used for chicken: ‘meat and skin, raw’; USDA National Nutrient Database terms used for pork: ‘composite of
trimmed leg, loin, shoulder, and spareribs, separable lean and fat, raw’ and ‘fresh, variety meats and by-products, mechanically separated, raw’.

Smil [23] (also cited in [20, 24]) provided protein and
calorie efficiency for farmed carp, however, Smil does
not provide methods or reference source data, making it
impossible to independenly reproduce the calculations
or update estimates as newer data becomes available.
Given the pressing challenges of limited resources and
rising global demand for animal products, it is criti-
cal to know which aquaculture species most efficiently
retain protein and calories in feed, and how aquatic
species compare to livestock. Our study fills this critical
research gap.
2. Methods
For this study, we calculated the protein and calo-
rie retention typical of commercial production for
several farmed aquatic and terrestrial animals by
developing equations and collecting data necessary
for filling in each variable. We included nine major
aquaculture species: common carp (Cyprinus carpio),
grass carp (Ctenopharyngodon idella), channel catfish
(Ictalurus punctatus), pangas catfish (Pangasius pan-
gasius), Atlantic salmon (Salmo salar), rainbow trout
(Oncorhynchus mykiss), giant tiger prawn (Penaeus
monodon), whiteleg shrimp (Litopenaeus vannamei),
tilapia (Oreochromis niloticus and other cichlids); and
three livestock groups (cattle raised for beef, pigs, and
chickens raised for meat). The aquaculture species
included in the study comprised over half (57%) of
global production of fed aquaculture in 2012 [10, 25],
and the livestock are the top land animals produced for
meat in the US and globally [9, 26].
We collected data from numerous sources on FCRs,
feed composition, yield/edible portion, and nutritional
profiles of edible flesh (table 1), and using these data we
calculated protein and calorie retention using equations
1 and 2. (See the supplementary material for additional
details regarding species selection, data extraction, and
development of the retention equations.) The equa-
tions we developed are comparable to the ‘Nutrient
retention’ measure in table S1, but no specific equation
was supplied in the paper that described the measure
[22] or other studies that calculated protein and calo-
rie retention for livestock. We focused on collecting
data that reflects intensive/commercial production, and
focused on top-producing countries where possible.
For example, we searched for feed content information
for complete, commercially available feeds and not sup-
plementary feeds used in extensive or semi-intensive
production settings. Mean retention values and stan-
dard deviations were calculated using Excel (Microsoft
Corp., Redmond, WA) and Crystal Ball (Oracle Corp.,
Redwood Shores, CA). For each species, two types of
simulations were run: protein retention and calorie
retention. The retention equation and values col-
lected for each variable were entered into the software.
A Monte Carlo simulation was run for each reten-
tion/species combination using custom fit distribution
settings, meaning the values were not expected to fit
a specific probability distribution. Each Monte Carlo
simulation ran 5000 trials; each trial randomly sampled
the values entered. Calculating the retention means and
standard deviations in this way allowed for inclusion of
multiple values and provided a range of likely retention
values based on all of the data we collected.

3
Environ. Res. Lett. 13 (2018) 024017

Giant tiger prawn

Common carp
Pangas catfish

Tilapia
Grass carp
Channel catfish
Whiteleg shrimp

Rainbow trout

Atlantic salmon
Aquaculture weighted avg.
Beef cattle
Pigs

Chicken

0 5% 10% 15% 20% 25% 30% 35% 40%

Retention
Calorie Retention Protein Retention

Figure 2. Protein and calorie retention for selected aquatic and terrestrial farmed animal species. Dots represent sample means and
bars represent standard deviations. Higher values indicate more efficient retention.

Equation (1). Protein retention of selected aquatic
and terrestrial animals.
Protein retention =
(g protein in edible portion)
(g protein in feed)
(edible portion) (g protein per 100 g of edible portion)
= (FCR) (g protein per 100 g feed)
Equation (2). Calorie retention of selected aquatic
and terrestrial animals.
(calories in edible portion)
Calorie retention = (calories in feed)
(edible portion) (calories per 100g edible portion)
= els. Similar to above, chicken and Atlantic salmon have
(FCR) (calories per 100 g feed)
3. Results
Based on global production levels for each aquatic
species (i.e. a weighted average), we estimate that for
every 100 g of protein in aquaculture feed for these nine
species/species groups, 19 g are available in the human
food supply (19% retention), and for every 100 kcal
in aquaculture feed, 10 kcal enter the human food
supply (10% retention) (figure 2). Protein and calorie
retention values for aquatic and terrestrial species were
similar. Protein retention means ranged from 14%–
28% for the nine aquatic species, and 13%–37% for
livestock. Calorie retention means ranged from 6%–
25% for the aquatic species, and 7%–27% for livestock.
Chickens performed best for both protein and calorie
retention, followed by Atlantic salmon.
The factors that drive protein retention are the
FCR, concentration of protein in feed, and edible por-
tion. There is little variation in protein levels in edible
flesh among aquatic and terrestrial species. Aquatic
species have FCRs similar to chickens and lower than
pigs and cattle, but require higher levels of protein in
their feed compared to livestock. For example, the
relatively high mean protein retention for Atlantic
salmon (28%) is due to a low FCR (1.2–1.5) and
high edible portion (0.58–0.88); these factors offset the
high levels of protein in Atlantic salmon feed (35.5%–
44%). Chicken has the highest mean protein retention
(37%), due to a low FCR (1.9), low feed protein
level (18%–23%), and high edible portion (0.70–.78)
(table 1).
For calorie retention, there is more variation in
calories in edible flesh (compared to the above-
described variation in protein in flesh) and less variation
in feed calories by species compared to feed protein lev-
the highest mean calorie retention: 27 and 25%, respec-
tively. Pigs have an FCR (3.9) that is less efficient than
chicken and aquatic species, but high calories in edible
flesh (211–304 kcal per 100 g) and the high edible por-
tion (0.68–0.76) improves pig calorie retention (16%).
Giant tiger prawn and tilapia have the lowest mean
calorie retention for aquaculture, 6% and 7%, respec-
tively. These values are driven by low calorie content in
edible flesh and low edible portions.
4. Discussion and conclusions
A limitation of relying solely on FCRs to assess effi-
ciency becomes apparent when comparing FCRs and
retention values (figures 1 and 2). If FCRs are a good
predictor of nutrients retained in outputs, calorie and
protein retention for aquatic species would be similar
to, or higher than, chicken. Instead, when using the pro-
tein and calorie retention measures as derived above,
the values for aquatic species are, with the exception
of Atlantic salmon, more similar to retention values
for pigs and cattle. Therefore, aquatic species, taken
together, have little or no efficiency benefit over live-
stock when assessed on the basis of these alternative

4
 Environ. Res. Lett. 13 (2018) 024017
Table 2. Mean FCR and retention values in the current study and past research.a

Current study Cassidy et al (2013) Shepon et al (2016)

FCRb Feed/edible Protein Calorie FCR Feed/edible Protein Calorie FCR Feed/edible Protein Calorie
weight retention retention weight retention retention weight retention retention

Common carp 1.7 3.78 0.15 0.09

Grass carp 1.7 3.78 0.18 0.09
5

Channel catfish 1.4 2.87 0.18 0.11

Pangas catfish 1.4 3.02 0.17 0.09
Atlantic salmon 1.3 1.77 0.28 0.25
Rainbow trout 1.3 2.14 0.22 0.16
Giant tiger prawn 1.7 4.25 0.14 0.06
Whiteleg shrimp 1.7 2.68 0.22 0.09
Tilapia 1.7 4.17 0.18 0.07
Aquatic weighted average 1.6 3.08 0.19 0.10
Beef cattle 8.0 14.0 0.13 0.07 12.7 21.2 0.05 0.03 14 36 0.03 0.03
Pigs 3.9 5.34 0.21 0.16 6.5 9.3 0.10 0.10 3.1 6.0 0.09 0.09
Chicken 1.9 2.57 0.37 0.27 2.5 3.3 0.40 0.12 1.9 4.2 0.21 0.13
Eggs 0.35 0.22 0.31 0.17
Dairy 0.43 0.40 0.14 0.17
a Sources: Cassidy et al (2013) [7] and Shepon et al (2016) [21].
b FCRs are average values based on Tacon and Metian (2008) (aquatic species) [12]; Smil (2013) (livestock species) [13]; Shike (2013) (cattle) [14]; Zuidhof et al (2014) (chicken) [15]; and Rabobank Research (2015) (pigs) [16].
Environ. Res. Lett. 13 (2018) 024017
measures, which is the opposite of the result when
comparing FCRs.
Animal production involves complex biological,
social, and economic systems that vary by producer,
region, and within particular breeds or sub-species of
animals. Identifying differences by geography or cli-
mate was not possible in the current study. A strength
of this study is identifying the range of likely pro-
tein and calorie efficiencies using existing commercial
farming practices. Large standard deviations for some
species may reflect higher uncertainty due to data vari-
ance and/or more data availability for certain species.
Our aquaculture results are consistent with a previous
Atlantic salmon estimate (27% protein and 24% calorie
retention in Ytresoyl et al [22]) and lower than a protein
retention estimate for carps (30% in Smil [23]; we cal-
culated 15% for common carp and 18% for grass carp).
Importantly, we found that Atlantic salmon performs
much better than, and is therefore not representative
of, other aquatic species.
We calculated retention values for livestock instead
of using values from existing sources to ensure
standardized methods were used across aquatic and
terrestrial species. The work of Cassidy et al and She-
pon et al use overall feed use and production data,
therefore taking breeding animals and early mortalities
into account [7, 21]. These data were not consistently
available for aquaculture, so our results are based on
estimated inputs fed directly to meat-producing ani-
mals to produce a unit of edible flesh, and does not
account for breeding animals, wasted feed, or mortali-
ties. Therefore, our results may overestimate retention.
Not surprisingly, given the data used, our livestock
retention results are generally higher than previous
studies (table 2). The data used in this study for
meat-producing animals was more robust compared
to previous studies due to inclusion of multiple sources
of data for many variables and use of Monte Carlo
simulations for each retention/species combination to
provide likely values based on all data collected.
Another reason our protein and calorie retention
results may overestimate retention for aquaculture and
livestock is due to our exclusion of feed and nutrient
composition data for the earliest life stages. Detailed
information on feed content across lifestage was not
available for all species in the study. The final lifestage
(i.e. growout) involves the majority of feed consump-
tion and weight gain. Feed used in the earliest lifestages
are higher in protein (this is true for aquatic and ter-
restrial animals), but smaller amounts of feed are used.
Future research could refine retention calculations by
using feed information across multiple lifestages.
Adequate protein and calories, as well as micronu-
trient intake, are critical for avoiding malnutrition in
humans. Aquaculture expansion has been promoted
as part of a solution to provide animal protein more
efficiently, but in the case of fed aquaculture, the
implications of consuming farmed aquatic animals
that have higher requirements for protein in their diet
6
compared to terrestrial animals (including humans)
and relatively low retention of protein and calories
must be considered. Fed aquaculture requires more
highly nutrient-dense feed than livestock production
(i.e. higher in protein and calories), and results in a
loss of most protein (81%) and calories (90%) during
production. Animal products’ contribution to calories
in the human food supply is less critical compared to
protein and some micronutrients, but understanding
retention of calories in animal feed is important due to
the massive reduction during animal production and
the resulting impact on the overall supply of calories.
In addition, relationships between feed requirements
and affordability of final products should be explored.
Feed costs make up more than 50% of production
costs in aquaculture [29], and requirements for highly
nutrient-dense feeds likely influence prices of some
farmed seafood even though FCRs are low. Unfed
aquaculture, including aquatic plants and certain
shellfish, represents a critical opportunity to expand
production of highly nutritious human food with no
feed inputs.
Animal agriculture, including fed aquaculture,
relies on increasing amounts of grains, cereals, oilcakes,
and other staple crops for feed inputs as production
expands [11, 30]. Arable land used to grow feed crops
can compete with alternative land uses such as growing
crops directly for human consumption, which pro-
vides a more efficient transfer of calories and protein
[2, 31, 32]. While food security advocates may wish
for a shift in land use towards edible crops, the finan-
cial calculus for farmers’ crop planting decisions are
tied to complex social, political, and economic forces
that may be difficult to change. It is important to
note that animals’ diets can include human-indigestible
forages or by-products not considered to be human-
edible (e.g. distillers grains, bloodmeal), depending on
the species and production method, which reduces
competition for food between animal agriculture and
humans. Efforts are underway to develop and scale-up
production of insects as animal feed (and for human
consumption), but insects are not presently a major
component of feed.
Future research should explore potential impli-
cations and tradeoffs of increasing demand for
aquaculture feed that is higher in protein and calories,
as compared to livestock feed that is higher in starches.
For example, crops that are high in starch have higher
yields per hectare than crops high in protein due, in
part, to the energy cost of nitrogen fixing for legumes
[33]; average yields in tonnes per hectare for 2010–2014
were: maize 5.27, wheat 3.15, soy 2.50, and ground-
nuts 1.66 [34]. Soy and groundnuts are a significant
source of protein in aquaculture feeds and demand for
these ingredients is growing [11]. Therefore, global land
requirements for feed production could expand more
rapidly than current trends indicate and demand for
nitrogen fertilizer could grow more slowly. Other topics
for additional investigation include protein and calorie
Environ. Res. Lett. 13 (2018) 024017
retention in non-intensive production settings, strate-
gies aquaculture producers can employ to improve
protein and calorie retention, and retention of marine
omega-3 fatty acids and micronutrients (e.g. iron) in
farmed seafood. Current efforts to reduce FCRs in
aquaculture and livestock production focus on genetic
improvements through breeding and genetic engi-
neering, development of nutritionally superior feeds
and supplements, identification and implementation
of improved husbandry practices including ideal envi-
ronmental conditions for faster animal growth, and
overcoming cost and other barriers to increase pro-
ducer access to all of these developments. In addition
to FCR, researchers should explore the impact of these
changes on protein and calorie retention. For example,
it is possible that providing feed higher in protein could
result in a more efficient FCR but less efficient protein
retention.
Our results reveal a different ranking in terms of
efficiency of farmed animal species when measured
by FCR versus protein and calorie retention. We do
not argue, however, that retention measures should
replace FCR as the major indicator of feed efficiency.
Instead, the best path forward is to use multiple mea-
sures to compare efficiency of various types of food
production, including animals and plants. Discussions
of sustainable food systems should be informed by a
combination of factors including FCRs and nutrient
retention, and also environmental footprint measures
including resource use (e.g. land, water, fertilizer),
greenhouse gas emissions, and negative externalities
including biodiversity loss and water pollution. To
facilitate uptake of retention measures by researchers
and other stakeholders, we provide our data sources
and equations, which can be refined and applied to
additional species, settings, and nutrients.
Growing evidence supports dietary shifts toward
more efficient types of animal protein and plant-based
foods [2, 4, 6, 7, 21]. In order to inform policy change
and other interventions, there is a need to better under-
stand the flow and loss of nutrients in the global food
system, especially for fed aquaculture, and the impli-
cations for resource use and global food security. Our
results show that protein and calorie retention in aqua-
culture varies by species, is lower than chickens, and
similar to pig and cattle production, despite lower
FCRs. This research demonstrates the importance of
assessing animal production efficiency using multiple
measures.
Acknowledgments
Support for J P F, N A M, D C L, and M C M was pro-
vided by the Johns Hopkins Center for a Livable Future
(CLF) with a gift from the GRACE Communications
Foundation. The authors thank Brent Kim, Roni Neff,
Shawn McKenzie, Jim Yager, and Leo Horrigan at CLF
for providing helpful feedback on the manuscript.
7
ORCID iDs
Jillian P Fry https://orcid.org/0000-0002-5836-9076
Nicholas A Mailloux https://orcid.org/0000-0002-
4041-3665
David C Love https://orcid.org/0000-0002-2606-
8623
References
[1] Rockström J et al 2009 A safe operating space for humanity
Nature 461 472–5
[2] Foley J A et al 2011 Solutions for a cultivated planet Nature
478 337–42
[3] Tilman D et al 2011 Global food demand and the sustainable
intensification of agriculture Proc. Natl Acad. Sci. USA 108
20260–4
[4] Wirsenius S, Azar C and Berndes G 2010 How much land is
needed for global food production under scenarios of dietary
changes and livestock productivity increases in 2030? Agric.
Syst. 103 621–38
[5] Ran Y et al 2016 Assessing water resource use in
livestock production: a review of methods Livest. Sci. 187
68–79
[6] Tilman D and Clark M 2014 Global diets link environmental
sustainability and human health Nature 515 518–22
[7] Cassidy E S et al 2013 Redefining agricultural yields: from
tonnes to people nourished per hectare Environ. Res. Lett. 8
034015
[8] UN Food and Agriculture Organization 2016 The State of
World Fisheries and Aquaculture (Rome: FAO) (Accessed: 9
November 2017) (www.fao.org/3/a-i5555e.pdf)
[9] United States Department of Agriculture 2017 Livestock and
Poultry: World Markets and Trade (Accessed: 9 November
2017) (https://apps.fas.usda.gov/psdonline/circulars/
livestock_poultry.pdf)
[10] Tacon A G J and Metian M 2015 Feed matters: satisfying
the feed demand of aquaculture Rev. Fish. Sci. Aquac. 23
1–10
[11] Fry J P et al 2016 Environmental health impacts of feeding
crops to farmed fish Environ. Int. 91 201–14
[12] Tacon A G J and Metian M 2008 Global overview on the use
of fish meal and fish oil in industrially compounded aquafeeds:
trends and future prospects Aquaculture 285 146–58
[13] Smil V 2013 Should we eat meat? Evolution and Consequences
of Modern Carnivory (Chichester: Wiley-Blackwell)
[14] Shike D W 2013 Beef Cattle Feed Efficiency (Accessed: 2
November 2017) (http://lib.dr.iastate.edu/cgi/viewcontent.
cgi?article=1027&context=driftlessconference)
[15] Zuidhof M J et al 2014 Growth, efficiency, and yield of
commercial broilers from 1957, 1978, and 2005 Poult. Sci. 93
2970–82
[16] Rabobank Food and Agribusiness Research 2015 Pigs Might
Fly: Peak Pork Production Potential (Accessed: 2 November
2017) (https://research.rabobank.com/far/en/sectors/farm-
inputs/Pigs_Might_Fly.html)
[17] Naylor R L et al 2009 Feeding aquaculture in an era of finite
resources Proc. Natl Acad. Sci. USA 106 15103–10
[18] Torrissen O et al 2011 Atlantic Salmon (Salmo salar): the
‘super-chicken’ of the Sea? Rev. Fish. Sci. 19 257–78
[19] Béné C et al 2005 Feeding 9 billion by 2050—putting fish back
on the menu Food Secur. 7 261–74
[20] High Level Panel of Experts (HLPE) on Food Security and
Nutrition 2014 Sustainable Fisheries and Aquaculture for Food
Security and Nutrition (Rome: FAO) (Accessed: 24 January
2017) (www.fao.org/3/a-i3844e.pdf)
[21] Shepon A et al 2016 Energy and protein feed-to-food
conversion efficiencies in the US and potential food
security gains from dietary changes Environ. Res. Lett. 11
105002
Environ. Res. Lett. 13 (2018) 024017
[22] Ytrestøyl T, Aas T S and Åsgård T 2015 Utilisation of feed
resources in production of Atlantic salmon (Salmo salar) in
Norway Aquaculture 448 365–74
[23] Smil V 2002 Nitrogen and food production: proteins for
human diets Ambio 31 126–31
[24] Hall S et al 2011 Blue Frontiers: Managing the Environmental
Costs of Aquaculture (Penang: FAO) (Accessed: 3 February
2017) (www.conservation.org/publications/documents/
BlueFrontiers_aquaculture_report.pdf)
[25] FAO, 2017 Fishery Statistical Collections 2017 (www.fao.org/
fishery/statistics/global-production/en)
[26] US Department of Agriculture Economic Research Service
2016 Livestock and Meat Domestic Data (Accessed: 25
January 2017) (www.ers.usda.gov/data-products/livestock-
meat-domestic-data/livestockmeat-domestic-data/#All-
meat-statistics)
[27] USDA National Agricultural Library 2016 National Nutrient
Database for Standard Reference Release 28 (https://ndb.
nal.usda.gov)
[28] Seafood Health Facts Pangasius (www.seafoodhealthfacts.
org/description-top-commercial-seafood-items/pangasius)
[29] Rana K J, Siriwardena S and Hasan M R 2009 Impact of Rising
Feed Ingredient Prices on Aquafeeds and Aquaculture
Production (Rome: FAO) (Accessed: 25 May 2017)
(www.fao.org/docrep/012/i1143e/i1143e00.htm)
[30] Alexandratos N and Bruinsma J 2012 World agriculture
towards 2030/2050: The 2012 revision 12-3 (Rome: FAO)
(Accessed: 29 June 2016) (www.fao.org/economic/esa)
[31] Gephart J A et al 2016 The environmental cost of subsistence:
optimizing diets to minimize footprints Sci. Total Environ. 553
120–7
[32] Davis K et al 2016 Meeting future food demand with
current agricultural resources Glob. Environ. Change 39
125–32
[33] Bues A et al 2013 The Environmental Role of Protein Crops in
the New Common Agricultural Policy (Brussels: European
Union) (Accessed: 17 May 2017) (www.europarl.europa.eu/
RegData/etudes/etudes/join/2013/495856/IPOL-AGRI_ET
(2013)495856_EN.pdf)
[34] OECD-FAO 2016 Agricultural Outlook 2016–2025 (Paris:
OECD Publishing) (http://stats.oecd.org/index.aspx?
queryid=71240)